The comparative morphology of the musculature controlling the pectoral free rays in scorpaenoid fishes.

Free rays are ventral pectoral fin rays (lepidotrichia) that are free of the pectoral fin webbing. They are some of the most striking adaptations of benthic fishes. Free rays are used for specialized behaviors such as digging, walking or crawling along the sea bottom. Studies of pectoral free rays have focused on a small number of species, most notably the searobins (Family Triglidae). Previous research on the morphology of the free rays has emphasized their functional novelty. We hypothesize that the more extreme specializations of the pectoral free rays in searobins are not precisely novel, but are part of a broader range of morphological specializations that are associated with the pectoral free rays in suborder Scorpaenoidei. We perform a detailed comparative description of the intrinsic musculature and osteology of the pectoral free rays in three families of scorpaenoid fishes: Hoplichthyidae, Triglidae, and Synanceiidae. These families vary in the number of pectoral free rays and the degree of morphological specialization of those rays. As part of our comparative analysis, we propose significant revisions to earlier descriptions of both the identity and function of the musculature associated with the pectoral free rays. We focus particularly on the specialized adductors that are important for walking behaviors. Our emphasis on the homology of these features provides important morphological and evolutionary context for understanding the evolution and function of free rays within Scorpaenoidei and other groups.

Fin ray patterns at the fin-to-limb transition.

The fin-to-limb transition was marked by the origin of digits and the loss of dermal fin rays. Paleontological research into this transformation has focused on the evolution of the endoskeleton, with little attention paid to fin ray structure and function. To address this knowledge gap, we study the dermal rays of the pectoral fins of 3 key tetrapodomorph taxa-Sauripterus taylori (Rhizodontida), Eusthenopteron foordi (Tristichopteridae), and Tiktaalik roseae (Elpistostegalia)-using computed tomography. These data show several trends in the lineage leading to digited forms, including the consolidation of fin rays (e.g., reduced segmentation and branching), reduction of the fin web, and unexpectedly, the evolution of asymmetry between dorsal and ventral hemitrichia. In Eusthenopteron, dorsal rays cover the preaxial endoskeleton slightly more than ventral rays. In Tiktaalik, dorsal rays fully cover the third and fourth mesomeres, while ventral rays are restricted distal to these elements, suggesting the presence of ventralized musculature at the fin tip analogous to a fleshy "palm." Asymmetry is also observed in cross-sectional areas of dorsal and ventral rays. Eusthenopteron dorsal rays are slightly larger than ventral rays; by contrast, Tiktaalik dorsal rays can be several times larger than ventral rays, and degree of asymmetry appears to be greater at larger sizes. Analysis of extant osteichthyans suggests that cross-sectional asymmetry in the dermal rays of paired fins is plesiomorphic to crown group osteichthyans. The evolution of dermal rays in crownward stem tetrapods reflects adaptation for a fin-supported elevated posture and resistance to substrate-based loading prior to the origin of digits.

Variation in flexural stiffness of the lepidotrichia within and among the soft fins of yellow perch under different preservation techniques.

Although the ray-finned fishes are named for their bony, segmented lepidotrichia (fin rays), we are only beginning to understand the morphological and functional diversity of this key vertebrate structure. Fin rays support the fin web, and their material properties help define the function of the entire fin. Many earlier studies of fin ray morphology and function have focused on isolated rays, or on rays from only one or two fins. At the same time, relatively little is known about how different preservation techniques affect the material properties of many vertebrate structures, including fin rays. Here, we use three-point bending tests to examine intra- and inter-fin variation in the flexural stiffness of fin rays from yellow perch, Perca flavescens. We sampled fin rays from individuals that were assigned to one of three preservation treatments: fresh, frozen, and preserved with formalin. The flexural stiffness of the fin rays varied within and among fins. Pelvic-fin rays were the stiffest, and pectoral fin rays the least stiff. The fin rays of the dorsal, anal, and caudal fins all had similar stiffness values, which were intermediate relative to those from the paired fins. The flexural stiffness of the fin rays was higher in rays that were at the leading edge of the fin. This variation in flexural stiffness was associated with variation in joint density and the relative length of the unsegmented proximal base of the fin rays. There was no significant difference in flexural stiffness between fresh and frozen specimens. In specimens preserved with formalin, there is a small but significant effect on stiffness in smaller fin rays.

Functional implications of morphological specializations among the pectoral fin rays of the benthic longhorn sculpin.

Fin ray structure in ray-finned fishes (Actinopterygii) largely defines fin function. Fin rays convert the muscle activity at the base of the fin to shape changes throughout the external fin web. Despite their critical functional significance, very little is known about the relationship between form and function in this key vertebrate structure. In this study we demonstrate that morphological specializations of the pectoral fin rays of the benthic longhorn sculpin (Myoxocephalus octodecimspinosus) have specific functional consequences both within and among individual rays. The fin rays of longhorn sculpin have an elongate unjointed region with a cylindrical shape in cross-section proximally, and are jointed with a crescent-shaped cross-section distally. Variation in the relative length of the proximal versus distal regions affects the location of maximum curvature as well as the mean curvature along the length of individual rays. We experimentally manipulated fin rays to mimic the differential muscle activity that generates curvature of fin rays in living animals. We found that the shape of the fin rays in cross-section affects their curvature. Among fin rays, the most ventral fin rays with relatively longer proximal unjointed regions have a more distal location of maximum curvature. These ventral rays also have higher mean curvature, likely because of a combination of features including the cross-sectional shape, area and diameter of the distal segments as well as their relative size and number, which were not examined in detail here. Because these rays are used routinely for substrate contact, this higher curvature could contribute to increased flexibility for substrate contact behaviors such as clinging or gripping the substrate. These morphological and functional differences among fin rays are correlated with the functional regionalization of the fin. Specifically, the ventral fin rays that are used during substrate contact are more stiff proximally and more highly curved distally than the pectoral rays in the dorsal region, which are longer and used during slow swimming. This study highlights the importance of examining morphological and functional variation both within and among complex structures such as fin rays.

Functional implications of variation in pectoral fin ray morphology between fishes with different patterns of pectoral fin use.

In this study, I compare the morphology from the pectoral fin rays from the benthic longhorn sculpin (Myoxocephalus octodecimspinosus) to those from a species that does not use its fins for substrate contact, the yellow perch (Perca flavescens). I use CT scanning technology to compare the shape and structure of the paired hemitrichia that make up the pectoral fin rays between these species. I found that the structure of hemitrichia of the fin rays in yellow perch is consistent with previous descriptions for pelagic fishes. They are almost completely segmented, have a crescent shape in cross section, and are branched distally. In contrast, longhorn sculpin hemitrichia exhibit morphological regionalization along the proximo-distal length of the ray. The most proximal 20-50% of the length of the hemitrichia is unsegmented and cylindrical in cross section. Distally, the fin rays of longhorn sculpin are segmented and crescent-shaped but do not branch. I measured the second moment of area of the hemitrichia at distances of 10%, 30%, 50%, and 70% distance along the length of the fin rays. The cylindrical regions of the sculpin hemitrichia had a higher second moment of area than the crescent-shaped regions in either species. I hypothesize that that this regionalization of individual fin rays provides resistance to bending proximally and flexibility distally, features that may be useful during substrate contact. This combination of an elongate, unsegmented proximal region and segmented distal region in fin rays has not yet been described among extant ray-finned fishes. However, this structure is reminiscent of that of the elongate cylindrical region found in the fossil sarcopterygian fish Eusthenopteron.